Selected Publications

@article{eusebi_gene_2021,
title = {Gene expression profiles underlying aggressive behavior in the prefrontal cortex of cattle},
volume = {22},
issn = {1471-2164},
url = {https://doi.org/10.1186/s12864-021-07505-5},
doi = {10.1186/s12864-021-07505-5},
abstract = {Aggressive behavior is an ancient and conserved trait, habitual for most animals in order to eat, protect themselves, compete for mating and defend their territories. Genetic factors have been shown to play an important role in the development of aggression both in animals and humans, displaying moderate to high heritability estimates. Although such types of behaviors have been studied in different animal models, the molecular architecture of aggressiveness remains poorly understood. This study compared gene expression profiles of 16 prefrontal cortex (PFC) samples from aggressive and non-aggressive cattle breeds: Lidia, selected for agonistic responses, and Wagyu, selected for tameness.},
number = {1},
urldate = {2021-05-31},
journal = {BMC Genomics},
author = {Eusebi, Paulina G. and Sevane, Natalia and O’Rourke, Thomas and Pizarro, Manuel and Boeckx, Cedric and Dunner, Susana},
month = apr,
year = {2021},
pages = {245},
}

@article{theofanopoulou_universal_2021,
title = {Universal nomenclature for oxytocin–vasotocin ligand and receptor families},
volume = {592},
copyright = {2021 The Author(s), under exclusive licence to Springer Nature Limited},
issn = {1476-4687},
url = {https://www.nature.com/articles/s41586-020-03040-7},
doi = {10.1038/s41586-020-03040-7},
abstract = {Oxytocin (OXT; hereafter OT) and arginine vasopressin or vasotocin (AVP or VT; hereafter VT) are neurotransmitter ligands that function through specific receptors to control diverse functions1,2. Here we performed genomic analyses on 35 species that span all major vertebrate lineages, including newly generated high-contiguity assemblies from the Vertebrate Genomes Project3,4. Our findings support the claim5 that OT (also known as OXT) and VT (also known as AVP) are adjacent paralogous genes that have resulted from a local duplication, which we infer was through DNA transposable elements near the origin of vertebrates and in which VT retained more of the parental sequence. We identified six major oxytocin–vasotocin receptors among vertebrates. We propose that all six of these receptors arose from a single receptor that was shared with the common ancestor of invertebrates, through a combination of whole-genome and large segmental duplications. We propose a universal nomenclature based on evolutionary relationships for the genes that encode these receptors, in which the genes are given the same orthologous names across vertebrates and paralogous names relative to each other. This nomenclature avoids confusion due to differential naming in the pre-genomic era and incomplete genome assemblies, furthers our understanding of the evolution of these genes, aids in the translation of findings across species and serves as a model for other gene families.},
language = {en},
number = {7856},
urldate = {2021-05-31},
journal = {Nature},
author = {Theofanopoulou, Constantina and Gedman, Gregory and Cahill, James A. and Boeckx, Cedric and Jarvis, Erich D.},
month = apr,
year = {2021},
pages = {747--755},
}

@article{orourke_capturing_2021,
title = {Capturing the {Effects} of {Domestication} on {Vocal} {Learning} {Complexity}},
issn = {13646613},
url = {https://linkinghub.elsevier.com/retrieve/pii/S1364661321001236},
doi = {10.1016/j.tics.2021.05.002},
language = {en},
urldate = {2021-05-20},
journal = {Trends in Cognitive Sciences},
author = {O'Rourke, Thomas and Martins, Pedro Tiago and Asano, Rie and Tachibana, Ryosuke O. and Okanoya, Kazuo and Boeckx, Cedric},
month = may,
year = {2021},
pages = {S1364661321001236},
}

@article{moriano_human-derived_2021,
type = {preprint},
title = {Human-derived alleles in SOST and RUNX2 3′{UTRs} cause differential regulation in a bone cell-line model},
url = {http://biorxiv.org/lookup/doi/10.1101/2021.04.21.440797},
abstract = {Abstract
The inquiry into the phenotypic features that set apart human species, such as a light, gracile skeleton and a rounded skull characteristic of
Homo sapiens
, can now benefit from the examination of ancient genomes. These have added a new layer of analysis allowing the identification of genetic differences between species like ours and our closest extinct relatives. Most of these genetic differences are non-coding changes with unknown functional consequences, and dissecting their putative regulatory effect remains challenging. Here we focus on the three prime untranslated regions (3’UTR), known to play a critical role in messenger RNA regulation and a plausible locus for divergent regulation between
Homo
species. We report a set of genes with derived 3’UTR changes in either the
Homo sapiens
or the Neanderthal/Denisovan lineages and experimentally evaluate the impact of 3’UTR variants in four genes:
E2F6
,
GLI3
,
RUNX2
and
SOST
. We performed a luciferase reporter assay in a bone cell-line model and found a statistically significant difference for the 3’UTR variants of
SOST
(
Homo sapiens
-derived) and
RUNX2
(Neanderthal/Denisovan-derived). The differential expression caused by these variants in our experimental model points to species differences in bone mineral density. Thus, this study adds insights into the functional effects of regulatory variants that emerged in recent human evolution.},
language = {en},
urldate = {2021-05-20},
institution = {Evolutionary Biology},
journal = {bioRxiv},
author = {Moriano, Juan and Martínez-Gil, Núria and Andirkó, Alejandro and Balcells, Susana and Grinberg, Daniel and Boeckx, Cedric},
month = apr,
year = {2021},
doi = {10.1101/2021.04.21.440797},
}

@article{buisan_distinct_2021,
title = {A distinct expression profile in the cerebellum and striatum for genes under selection within introgression deserts},
copyright = {© 2021, Posted by Cold Spring Harbor Laboratory. This pre-print is available under a Creative Commons License (Attribution-NonCommercial-NoDerivs 4.0 International), CC BY-NC-ND 4.0, as described at http://creativecommons.org/licenses/by-nc-nd/4.0/},
url = {https://www.biorxiv.org/content/10.1101/2021.03.26.437167v2},
doi = {10.1101/2021.03.26.437167},
abstract = {{\textless}h3{\textgreater}Abstract{\textless}/h3{\textgreater} {\textless}p{\textgreater}Analyses of ancient DNA from extinct hominins have provided unique insights into the complex evolutionary history of \textit{Homo sapiens}, intricately related to that of the Neanderthals and the Denisovans as revealed by several instances of admixture events. These analyses have also allowed the identification of introgression deserts: genomic regions in our species that are depleted of ‘archaic’ haplotypes. The presence of genes like \textit{FOXP2} in these deserts has been taken to be suggestive of brain-related functional differences between \textit{Homo} species. Here, we seek a deeper characterization of these regions, taking into account signals of positive selection in our lineage. Analyzing publicly available transcriptomic data from the human brain at different developmental stages, we found that structures outside the cerebral neocortex, and especially the cerebellum and the striatum at prenatal stages, show the most divergent transcriptomic profiles when considering genes under positive selection within introgression deserts.{\textless}/p{\textgreater}},
language = {en},
urldate = {2021-05-20},
journal = {bioRxiv},
author = {Buisan, Raül and Moriano, Juan and Andirkó, Alejandro and Boeckx, Cedric},
month = apr,
year = {2021},
pages = {2021.03.26.437167},
}

@article{andirko_fine-grained_2021,
title = {Fine-grained temporal mapping of derived high-frequency variants supports the mosaic nature of the evolution of {Homo} sapiens},
copyright = {© 2021, Posted by Cold Spring Harbor Laboratory. This pre-print is available under a Creative Commons License (Attribution-NonCommercial-NoDerivs 4.0 International), CC BY-NC-ND 4.0, as described at http://creativecommons.org/licenses/by-nc-nd/4.0/},
url = {https://www.biorxiv.org/content/10.1101/2021.01.22.427608v1},
doi = {10.1101/2021.01.22.427608},
abstract = {{\textless}h3{\textgreater}ABSTRACT{\textless}/h3{\textgreater} {\textless}p{\textgreater}As our knowledge about the history of the \textit{Homo sapiens} lineage becomes increasingly complex, large-scale estimations of the time of emergence of derived variants become essential to be able to offer more precise answers to time-sensitive hypotheses concerning human evolution. Using an open repository of genetic variant age estimations recently made available, we offer here a temporal evaluation of various evolutionarily relevant datasets, such as \textit{Homo sapiens}-specific variants, high-frequency variants found in genetic windows under positive selection, introgressed variants from extinct human species, as well as putative regulatory variants in various brain regions. We find a recurrent bimodal distribution of high-frequency variants, but also evidence for specific enrichments of gene categories in various time windows, which brings into prominence the 300-500k time slice. We also find evidence for very early mutations impacting the facial phenotype, and much more recent molecular events linked to specific brain regions such as the cerebellum or the precuneus. Additionally, we present a case study of an evolutionarily relevant gene, \textit{BAZ1B}, and its targets, to emphasize the importance of applying temporal data to specific evolutionary questions. Overall, we present a unique resource that informs and complements our previous knowledge of \textit{Homo sapiens} evolution using publicly available data, and reinforce the case for the mosaic, temporally very extended nature of the evolutionary trajectory of our species.{\textless}/p{\textgreater}},
language = {en},
urldate = {2021-05-20},
journal = {bioRxiv},
author = {Andirkó, Alejandro and Moriano, Juan and Vitriolo, Alessandro and Kuhlwilm, Martin and Testa, Giuseppe and Boeckx, Cedric},
month = jan,
year = {2021},
pages = {2021.01.22.427608},
}

@article{ruland_agent-based_2020,
title = {An {Agent}-based model of the gradual emergence of modern linguistic complexity},
url = {http://biorxiv.org/lookup/doi/10.1101/2020.11.12.380683},
language = {en},
urldate = {2020-11-17},
author = {Ruland, Marcel and Andirkó, Alejandro and Romanowska, Iza and Boeckx, Cedric},
month = nov,
year = {2020},
doi = {10.1101/2020.11.12.380683},
journal = {BioRxiv}
}

@inproceedings{evolang13_32,
author = "de Boer, B. and Thompson, B. and Ravignani, A. and Boeckx, C.",
title = "Analysis of Mutation and Fixation for Language",
year = "2020",
booktitle = "The Evolution of Language: Proceedings of the 13th International Conference (EvoLang13)",
editor = "Ravignani, A. and Barbieri, C. and Martins, M. and Flaherty, M. and Jadoul, Y. and Lattenkamp, E. and Little, H. and Mudd, K. and Verhoef, T.",
doi = "10.17617/2.3190925",
url = "http://brussels.evolang.org/proceedings/paper.html?nr=32",
}

@inproceedings{evolang13_81,
author = "Moriano Palacios, J. and Andirko, A. and Boeckx, C.",
title = "Genetic Networks Under Human-Specific Regulation Reveals Clues About the Evolution of the Modern Language-Ready Brain",
year = "2020",
booktitle = "The Evolution of Language: Proceedings of the 13th International Conference (EvoLang13)",
editor = "Ravignani, A. and Barbieri, C. and Martins, M. and Flaherty, M. and Jadoul, Y. and Lattenkamp, E. and Little, H. and Mudd, K. and Verhoef, T.",
doi = "10.17617/2.3190925",
url = "http://brussels.evolang.org/proceedings/paper.html?nr=81",
}

@inproceedings{evolang13_90,
author = "O'Rourke, T. and Boeckx, C.",
title = "Glutamate Receptors Implicated in (Self-)Domestication Regulate Dopaminergic Signaling in Striatal Vocal-Learning Pathways",
year = "2020",
booktitle = "The Evolution of Language: Proceedings of the 13th International Conference (EvoLang13)",
editor = "Ravignani, A. and Barbieri, C. and Martins, M. and Flaherty, M. and Jadoul, Y. and Lattenkamp, E. and Little, H. and Mudd, K. and Verhoef, T.",
doi = "10.17617/2.3190925",
url = "http://brussels.evolang.org/proceedings/paper.html?nr=90",
}

@inproceedings{martins_boeckx_2020,
author = {Martins, Pedro Tiago and Boeckx, Cedric"},
title = {Specifying the "Vocal" in Vocal Learning},
year = {2020},
booktitle = {The Evolution of Language: Proceedings of the 13th International Conference (EvoLang13)},
editor = {Ravignani, Andrea and Barbieri, Chiara and Martins, Maurício and Flaherty, Molly and Jadoul, Yannick and Lattenkamp, Ella and Little, Hannah and Mudd, Katie and Verhoef, Tessa},
doi = {10.17617/2.3190925},
url = {http://brussels.evolang.org/proceedings/paper.html?nr=150},
}

@inproceedings{silventeifont_2020,
author = {Silvente i Font, Sara and Martins, Pedro Tiago and Boeckx, Cedric},
title = {Do Life Histories Shape Vocal Production Learning? A Bird-Based Approach},
year = {2020},
booktitle = {The Evolution of Language: Proceedings of the 13th International Conference (EvoLang13)},
editor = {Ravignani, Andrea and Barbieri, Chiara and Martins, Maurício and Flaherty, Molly and Jadoul, Yannick and Lattenkamp, Ella and Little, Hannah and Mudd, Katie and Verhoef, Tessa},
doi = {10.17617/2.3190925},
url = {http://brussels.evolang.org/proceedings/paper.html?nr=147},
}

@article{Moriano_2020,
doi = {10.1186/s12864-020-6706-x},
year = {2020},
month = {apr},
publisher = {Springer Science and Business Media {LLC}},
volume = {21},
number = {1},
author = {Juan Moriano and Boeckx, Cedric},
abstract = {Recent paleogenomic studies have highlighted a very small set of proteins carrying modern human-specific missense changes in comparison to our closest extinct relatives. Despite being frequently alluded to as highly relevant, species-specific differences in regulatory regions remain understudied. Here, we integrate data from paleogenomics, chromatin modification and physical interaction, and single-cell gene expression of neural progenitor cells to identify derived regulatory changes in the modern human lineage in comparison to Neanderthals/Denisovans. We report a set of genes whose enhancers and/or promoters harbor modern human single nucleotide changes and are active at early stages of cortical development.},
title = {Modern human changes in regulatory regions implicated in cortical development},
journal = {{BMC} Genomics}
}

@article{martins2020,
author = {Martins, Pedro Tiago and Boeckx, Cedric},
title = {Vocal learning: Beyond the continuum},
journal = {PLoS Biology},
volume = {18},
number = {3},
pages = {e3000672},
year = {2020},
abstract = {Vocal learning is the ability to modify vocal output on the basis of experience. Traditionally, species have been classified as either displaying or lacking this ability. A recent proposal, the vocal learning continuum, recognizes the need to have a more nuanced view of this phenotype and abandon the yes–no dichotomy. However, it also limits vocal learning to production of novel calls through imitation, moreover subserved by a forebrain-to-phonatory-muscles circuit. We discuss its limitations regarding the characterization of vocal learning across species and argue for a more permissive view.},
doi = {https://doi.org/10.1371/journal.pbio.3000672}
}

@article{de_Boer_2020,
doi = {10.1038/s41598-019-57235-8},
url = {https://doi.org/10.1038%2Fs41598-019-57235-8},
year = {2020},
publisher = {Springer Science and Business Media {LLC}},
volume = {10},
number = {1},
author = {de Boer, Bart and Thompson, Bill and Ravignani, Andrea and Boeckx, Cedric},
title = {Evolutionary Dynamics Do Not Motivate a Single-Mutant Theory of Human Language},
abstract = {One of the most controversial hypotheses in cognitive science is the Chomskyan evolutionary conjecture that language arose instantaneously in humans through a single mutation. Here we analyze the evolutionary dynamics implied by this hypothesis, which has never been formalized before. The hypothesis supposes the emergence and fixation of a single mutant (capable of the syntactic operation Merge) during a narrow historical window as a result of frequency-independent selection under a huge fitness advantage in a population of an effective size no larger than ~15 000 individuals. We examine this proposal by combining diffusion analysis and extreme value theory to derive a probabilistic formulation of its dynamics. We find that although a macro-mutation is much more likely to go to fixation if it occurs, it is much more unlikely a priori than multiple mutations with smaller fitness effects. The most likely scenario is therefore one where a medium number of mutations with medium fitness effects accumulate. This precise analysis of the probability of mutations occurring and going to fixation has not been done previously in the context of the evolution of language. Our results cast doubt on any suggestion that evolutionary reasoning provides an independent rationale for a single-mutant theory of language.},
journal = {Scientific Reports}
}

@article{Orourke_2019_names,
title = "Names and their meanings: A dual-process account of proper-name encoding and retrieval",
journal = "Neuroscience & Biobehavioral Reviews",
volume = "108",
pages = "308 - 321",
year = "2020",
issn = "0149-7634",
doi = "https://doi.org/10.1016/j.neubiorev.2019.11.005",
url = "http://www.sciencedirect.com/science/article/pii/S0149763419302921",
author = "Thomas O’Rourke and Ruth de Diego Balaguer",
abstract = "The ability to pick out a unique entity with a proper name is an important component of human language. It has been a primary focus of research in the philosophy of language since the nineteenth century. Brain-based evidence has shed new light on this capacity, and an extensive literature indicates the involvement of distinct fronto-temporal and temporo-occipito-parietal association cortices in proper-name retrieval. However, comparatively few efforts have sought to explain how memory encoding processes lead to the later recruitment of these distinct regions at retrieval. Here, we provide a unified account of proper-name encoding and retrieval, reviewing evidence that socio-emotional and unitized encoding subserve the retrieval of proper names via anterior-temporal–prefrontal activations. Meanwhile, non-unitized item–item and item–context encoding support subsequent retrieval, largely dependent on the temporo-occipito-parietal cortex. We contend that this well-established divergence in encoding systems can explain how proper names are later retrieved from distinct neural structures. Furthermore, we explore how evidence reviewed here can inform a century-and-a-half-old debate about proper names and the meanings they pick out."
}

@article {Andirko_Boeckx_2020,
author = {Andirk{\'o}, Alejandro and Boeckx, Cedric},
title = {Modern human alleles differentially regulate gene expression in brain tissues: implications for brain evolution},
year = {2020},
doi = {10.1101/771816},
publisher = {Cold Spring Harbor Laboratory},
abstract = {The availability of high-coverage genomes of our extinct relatives, the Neanderthals and Denisovans, and the emergence of large, tissue-specific databases of modern human genetic variation, offer the possibility of probing the evolutionary trajectory of heterogenous structures of great interest, such as the brain. Here we cross two publicly available datasets, the GTEX cis-eQTL database (version 8) and an extended catalog of Homo sapiens specific alleles relative to the Neanderthal and Denisovan sequences to understand how nearly fixed Sapiens-derived alleles affect the regulation of gene expression across 15 structures. The list of variants obtained reveals enrichments in regions of the modern human genome showing putative signals of positive selection relative to archaic humans, points to associations with clinical conditions, and places the focus on specific structures such as the cerebellum and the Hypothalamus-Pituitary-Adrenal Gland axis. The directionality of regulation of these variants complements earlier findings about introgressed variants from archaics, and highlights the role of genes that deserve closer experimental attention.Competing Interest StatementThe authors have declared no competing interest.},
URL = {https://www.biorxiv.org/content/early/2020/04/21/771816},
journal = {bioRxiv}
}

@article{Kuhlwilm_2019,
doi = {10.1038/s41598-019-44877-x},
url = {https://doi.org/10.1038%2Fs41598-019-44877-x},
year = {2019},
volume = {9},
number = {1},
author = {Martin Kuhlwilm and Cedric Boeckx},
title = {A catalog of single nucleotide changes distinguishing modern humans from archaic hominins},
abstract = {Throughout the past decade, studying ancient genomes has provided unique insights into human prehistory, and differences between modern humans and other branches like Neanderthals can enrich our understanding of the molecular basis of unique modern human traits. Modern human variation and the interactions between different hominin lineages are now well studied, making it reasonable to go beyond fixed genetic changes and explore changes that are observed at high frequency in present-day humans. Here, we identify 571 genes with non-synonymous changes at high frequency. We suggest that molecular mechanisms in cell division and networks affecting cellular features of neurons were prominently modified by these changes. Complex phenotypes in brain growth trajectory and cognitive traits are likely influenced by these networks and other non-coding changes presented here. We propose that at least some of these changes contributed to uniquely human traits, and should be prioritized for experimental validation.},
journal = {{Scientific Reports}}
}

@article{martinsboeckx2019,
title = {Language evolution and complexity considerations: The no half-Merge fallacy},
author = {Martins, Pedro Tiago and Boeckx, Cedric},
year = {2019},
abstract = {Recently, prominent theoretical linguists have argued for an explicit scenario for the evolution of the human language capacity on the basis of its computational properties. Concretely, the simplicity of a minimalist formulation of the operation Merge, which allows humans to recursively compute hierarchical relations in language, has been used to promote a sudden-emergence, single-mutation scenario. In support of this view, Merge is said to be either fully present or fully absent: one cannot have half-Merge. On this basis, it is inferred that the emergence of our fully fledged language capacity had to be sudden. Thus, proponents of this view draw a parallelism between the formal complexity of the operation at the computational level and the number of evolutionary steps it must imply. Here, we examine this argument in detail and show that the jump from the atomicity of Merge to a single-mutation scenario is not valid and therefore cannot be used as justification for a theory of language evolution along those lines.},
journal = {PLoS Biology},
volume = {17},
number = {11},
pages = {e3000389},
doi = {https://doi.org/10.1371/journal.pbio.3000389}
}

@article {Zanella_etal_2019,
author = {Zanella, Matteo and Vitriolo, Alessandro and Andirko, Alejandro and Martins, Pedro Tiago and Sturm, Stefanie and O{\textquoteright}Rourke, Thomas and Laugsch, Magdalena and Malerba, Natascia and Skaros, Adrianos and Trattaro, Sebastiano and Germain, Pierre-Luc and Mihailovic, Marija and Merla, Giuseppe and Rada-Iglesias, Alvaro and Boeckx, Cedric and Testa, Giuseppe},
title = {Dosage analysis of the 7q11.23 Williams region identifies BAZ1B as a major human gene patterning the modern human face and underlying self-domestication},
volume = {5},
number = {12},
year = {2019},
doi = {10.1126/sciadv.aaw7908},
abstract = {We undertook a functional dissection of chromatin remodeler BAZ1B in neural crest (NC) stem cells (NCSCs) from a uniquely informative cohort of typical and atypical patients harboring 7q11.23 copy number variants. Our results reveal a key contribution of BAZ1B to NCSC in vitro induction and migration, coupled with a crucial involvement in NC-specific transcriptional circuits and distal regulation. By intersecting our experimental data with new paleogenetic analyses comparing modern and archaic humans, we found a modern-specific enrichment for regulatory changes both in BAZ1B and its experimentally defined downstream targets, thereby providing the first empirical validation of the human self-domestication hypothesis and positioning BAZ1B as a master regulator of the modern human face. In so doing, we provide experimental evidence that the craniofacial and cognitive/behavioral phenotypes caused by alterations of the Williams-Beuren syndrome critical region can serve as a powerful entry point into the evolution of the modern human face and prosociality.},
URL = {https://advances.sciencemag.org/content/5/12/eaaw7908},
journal = {Science Advances}
}

@inproceedings{martins_boeckx_17,
author = {Martins, Pedro Tiago and Boeckx, Cedric},
title = {Sound Production Learning and the Contiguum Hypothesis},
booktitle = {{Proceedings of Evolang XII}},
year = {2018},
url = {http://evolang.org/torun/proceedings/papertemplate.html?p=104},
doi = {10.12775/3991-1.069}
}

@article {TheoAndirkoBoeckx2018,
author = {Theofanopoulou, Constantina and Andirko, Alejandro and Boeckx, Cedric},
title = {Oxytocin and Vasopressin Receptor variants as a window onto the evolution of human prosociality},
year = {2018},
doi = {10.1101/460584},
abstract = {Modern humans{\textquoteright} lifestyle strongly depends on complex social skills like empathy, tolerance and cooperation. Variation in the oxytocin receptor (OXTR) and the arginine-vasopressin receptors (AVPR1A, AVPR1B genes) has been widely associated with diverse facets of social cognition, but the extent to which these variants may have contributed to the evolution of human prosociality remains to be elucidated. In this study, we compared the OXTR, AVPR1A and AVPR1B DNA sequences of modern humans to those of our closest extinct and extant relatives, and then clustered the variants we identified based on their distribution in the species studied. This clustering, along with the functional importance retrieved for each variant and their frequency in different modern-human populations, is then used to determine if any of the OXTR, AVPR1A and AVPR1B-variants might have had an impact at different evolutionary stages. We report a total of 29 SNPs, associated with phenotypic effects ranging from clearly pro-social to mixed or antisocial. Regarding modern human-specific alleles that could correlate with a shift towards prosociality in modern-humans, we highlight one allele in AVPR1A (rs11174811), found at high frequency and linked to prosocial phenotypes in modern humans, while the ancestral allele is associated with antisocial phenotypes. We also report three sites of putatively convergent changes between modern humans and bonobos (rs237897(A), rs2228485(G) and rs1042615(A)), and note the absence of such a convergent pattern between modern humans and chimpanzees. Finally, we observe the high concentration of {\textquoteright}modern human specific{\textquoteright} alleles in vasopressin receptors not paralleled in the oxytocin receptor.},
URL = {https://www.biorxiv.org/content/early/2018/11/04/460584},
eprint = {https://www.biorxiv.org/content/early/2018/11/04/460584.full.pdf},
journal = {{bioRxiv}}
}

@article{Orourke_2019_glutamate,
title = "Glutamate receptors in domestication and modern human evolution",
journal = "Neuroscience & Biobehavioral Reviews",
volume = "108",
pages = "341 - 357",
year = "2020",
issn = "0149-7634",
doi = "https://doi.org/10.1016/j.neubiorev.2019.10.004",
url = "http://www.sciencedirect.com/science/article/pii/S0149763419305433",
author = "Thomas O’Rourke and Cedric Boeckx",
abstract = "There has been a recent resurgence of interest in the hypothesis that anatomically modern humans and domesticated species have followed convergent evolutionary paths. Here, we review results from domestication and modern-human evolutionary studies in order to evaluate evidence for shared changes to neurotransmission across these species. We compare genomic and, where available, brain-expression differences across 488 neurotransmitter receptor genes in 14 domesticated species and modern humans relative to their wild and archaic counterparts. This analysis highlights prevalent changes to glutamate — most notably kainate and metabotropic — receptor genes. We review evidence for these genes’ expression and their respective receptor functions in the central nervous system, as well as phenotypes commonly associated with alterations to them. This evidence suggests an important role for kainate and metabotropic receptors in regulating hypothalamic–pituitary–adrenal axis excitation, and we provide a mechanistic account of their actions in attenuating the stress response. We assess the explanatory potential of such actions in contributing to the emergence of the (self-)domesticated phenotype, in particular to reduced reactive aggression."
}

@article {martins_etal2017,
author = {Martins, Pedro Tiago and Marí, Maties and Boeckx, Cedric},
title = {{SRGAP2} and the gradual evolution of the modern human language faculty},
year = {2018},
abstract = {In this paper we examine a new source of evidence that draws on data from archaic human genomes to support the hypothesis that vocal learning in Homo preceded the emergence of Anatomically Modern Humans. We build our claim on the evolutionary history of the SLIT-ROBO GTPase 2 gene (SRGAP2). The SLIT-ROBO molecular pathway has been shown to have an important role in the context of vocal learning. Though the relevance of the SRGAP2 gene duplication in the emergence of some aspect of language has not gone completely unnoticed, recent results now allow us to articulate a mechanistic hypothesis of its role in the context of axon guidance. Specifically, SRGAP2C, a duplication of SRGAP2 crucially also found in Neanderthals and Denisovans, but not in extant mammals, inhibits the ancestral SRGAP2A, which in turn modulates the axon guidance function of the SLIT-ROBO molecular pathway. This, we claim, could have contributed to the establishment of the critical cortico-laryngeal connection of the vocal learning circuit. Our conclusions support the idea that complex vocal learning could already have been part of the arsenal of some of our extinct ancestors.},
journal={{Journal of Language Evolution}},
pages = {lzx020},
doi = {10.1093/jole/lzx020}
}

@incollection{boeckx17langevol,
author = {Boeckx, Cedric},
year = {2017},
booktitle = {Evolution of Nervous Systems},
title = {Language evolution},
editor = {Kaas, J},
pages = {325--339},
publisher = {Elsevier},
address = {Oxford},
edition = {2},
volume = {4}
}

@article{theofanopoulou2016implications,
title={Implications of oxytocin in human linguistic cognition: from genome to phenome},
author={Theofanopoulou, Constantina},
journal={{Frontiers in Neuroscience}},
volume={10},
year={2016},
publisher={Frontiers Media SA},
doi = {10.3389/fnins.2016.00271}
}

@article{nunez2017large,
title={A large-scale study on the effects of sex on gray matter asymmetry},
author={N{\'u}{\~n}ez, Christian and Theofanopoulou, Constantina and Senior, Carl and Cambra, Maria Rosa and Usall, Judith and Stephan-Otto, Christian and Br{\'e}bion, Gildas},
journal={{Brain Structure and Function}},
volume = {223},
number = {1},
pages={183--193},
year={2017},
publisher={Springer},
doi = {10.1007/s00429-017-1481-4}
}

@article{theofanopoulou2017hypothesis,
title={A hypothesis on a role of oxytocin in the social mechanisms of speech and vocal learning},
author={Theofanopoulou, Constantina and Boeckx, Cedric and Jarvis, Erich D},
journal={{Proceedings of the Royal Society B}},
volume={284},
number={1861},
year={2017},
doi = {10.1098/rspb.2017.0988},
abstract = {Language acquisition in humans and song learning in songbirds naturally happen as a social learning experience, providing an excellent opportunity to reveal social motivation and reward mechanisms that boost sensorimotor learning. Our knowledge about the molecules and circuits that control these social mechanisms for vocal learning and language is limited. Here we propose a hypothesis of a role for oxytocin (OT) in the social motivation and evolution of vocal learning and language. Building upon existing evidence, we suggest specific neural pathways and mechanisms through which OT might modulate vocal learning circuits in specific developmental stages.}
}

@article{samuels17,
author = {Samuels, Bridget D and Martins, Pedro Tiago and Boeckx, Cedric},
year = {2017},
title = {言語知識の由来―強勢類型への進化的アプローチ― [Linguistic knowledge by descent: an evolutionary approach to stress typology]},
journal = {{音声研究 [Journal of the Phonetic Society of Japan]. 特集テーマ：音声/音韻から言語の化石を発掘する：進化言語学の最新の知見 [Special issue: Excavating phonetic/phonological fossils in language: current trends in evolutionary linguistics]}},
volume = {21},
number = {1},
pages = {71--78},
abstract = {We consider here several properties of phonological stress systems, including the midpoint pathology, an unattested pattern in which stress is confined to a word-medial syllable in short words but reverts to an edge-based window in longer words. Previous attempts have been made to rule out midpoint systems by eliminating the phonological constraints that yield them, or by alluding to difficulties in learning them. We suggest that a preference for representing word edges in memory and limits on subitization—evolutionarily older “fossil” abilities which are neither specific to humans nor to language — are sufficient to rule out the midpoint pathology. We take the same approach to motivate accentual window size and some left-right asymmetries observed in the typology of attested stress systems. This approach highlights the relevance of descent in accounting for human cognition, as well as the benefits that evolutionary thinking can bring to the study of language.},
url = {https://www.jstage.jst.go.jp/article/onseikenkyu/21/1/21_71/_pdf}
}

@article{reluprev,
Abstract = {As its title suggests, this is a biolinguistics book. Given the vast scope of this journal and the methodological and theoretical departures that the work under analysis might represent, I will start by making some very brief considerations, at the cost of not being able to provide an in-depth review of the book itself due to space restrictions, but with the advantage of providing readers with the right mindset: a fundamentally biolinguistic one, which hopefully enables a better understanding of Samuels' endeavor and its implications for those who are interested.},
Author = {Martins, Pedro Tiago},
Journal = {{Linguística - Revista de Estudos Linguísticos da Universidade do Porto}},
Pages = {209--216},
Title = {{Review of ``Phonological Architecture: A Biolinguistic Perspective'' (Bridget Samuels, 2011)}},
Volume = {7},
Year = {2012},
url = {http://ler.letras.up.pt/uploads/ficheiros/10752.pdf}
}

@incollection{theofanopoulou_thalamus,
author = {Theofanopoulou, Constantina and Boeckx, Cedric},
title = {The central role of the thalamus in language and cognition},
booktitle = {Advances in Biolinguistics: The Human Language Faculty and its Biological Basis},
editor = {Fujita, Koji and Boeckx, Cedric},
publisher = {Routledge},
address = {London},
year = {2016}
}

@article{theofanopoulou2015brain,
title={Brain asymmetry in the white matter making and globularity},
author={Theofanopoulou, Constantina},
journal={{Frontiers in Psychology}},
doi = {10.3389/fpsyg.2015.01355},
volume={6},
year={2015},
publisher={Frontiers Media SA}
}

@article{bird,
Author = {Martins, Pedro Tiago},
Journal = {Biolinguistics},
Pages = {97--107},
Title = {{The Bird is the Word: Review of ``Birdsong, Speech, Language: Exploring the Evolution of Mind and Brain'' (Ed. by Johan J. Bolhuis & Martin Everaert, 2013)}},
abstract = {The idea that birds might have something related to language that humans also seem to have has gone full circle: After the developments of linguistics and psychology during the 20th century put the ‘uniquely human’ in the center stage, with the help of failed or misled language experiments with animals, it now seems that perhaps birds have something to tell us after all. Even though the study of our closest cousins still very much dominates the understanding of our own biological and behavioral traits and tendencies, current, cutting-edge theories of language evolution now give a great deal of importance to the study of birds and their vocal abilities. It is not the case of course that scientists nowadays think that birds have ‘human language’ (they don’t, as the reader will also have concluded, if he has ever been around birds and tried to have a conversation). Instead, what has happened is that recent developments in various fields have made the study of birds a perfectly fine component of any serious approach to the unveiling of the nature of language.},
Volume = {8},
Year = {2014},
url = {http://www.biolinguistics.eu/index.php/biolinguistics/article/download/339/331}
}

@incollection{martinsetaladvances,
Address = {London},
Author = {Martins, Pedro Tiago and Leivada, Evelina and Benítez-Burraco, Antonio and Boeckx, Cedric},
Booktitle = {Advances in Biolinguistics: The Human Language Faculty and its Biological Basis},
Editor = {Fujita, Koji and Boeckx, Cedric},
Publisher = {Routledge},
Title = {Biological pluralism in service of biolinguistics},
Year = {2016},
pages = {153--169},
url = {https://www.routledge.com/Advances-in-Biolinguistics-The-Human-Language-Faculty-and-Its-Biological/Fujita-Boeckx/p/book/9781138891722},
abstract = {Excerpt: Upon close inspection, one concludes that the larger part of the issues that most generative linguistics work covers are philological in character, albeit through the use of sophisticated tools and notation.1 Thus, a very important (and unfortunate) realization when looking at the generative linguistics litera- ture is that its main premise (that language is a biological property of humans) does not entail, guide or constrain linguistic research in any meaningful way. In other words, if that premise were not held, the import of most linguistic work would remain largely unaffected, which is quite odd, for that premise is, again, one of the main tenets of generative linguistics. This apparent lack
interest in the biological half of biolinguistics is one of two problems regarding how investigations into the nature of the language faculty have been carried out. The second, related problem is the conception of biology itself that has served as the (rhetorical) backbone of some of these investigations. We will discuss the treatment that notions like novelty and variation have received in the linguistic literature, sometimes under the “biolinguistics” rubric, and offer some insights and counter-evidence from evolutionary biology, in favor of a biologically informed study of language.}
}

@incollection{martinsboeckxproblem,
author = {Martins, Pedro Tiago and Boeckx, Cedric},
title = {Language evolution: Insisting on making it a mystery or turning it into a problem?},
booktitle = {Papers dedicated to {Anne Reboul}},
editor = {Dupuy, Ludivine and Grabizna, Adrianna and Foudon, Nadège and Saint-Germier, Pierre},
publisher = {Institut des Sciences Cognitives/CNRS},
address = {Lyon},
pages = {1--10},
url = {http://reboul.isc.cnrs.fr/MartinsBoeckx.pdf},
year = {2016},
abstract = {In a recent, widely-read paper, Hauser et al. (2014) offer a rather negative view of the state of affairs in language evolution. More speciffcally, the authors believe that little to no progress has been made in the various relevant fields regarding the age-old questions of the origin and evolution of the human capacity for language. We beg to differ.}
}

@inproceedings{martins15apl,
author={Martins, Pedro Tiago},
year={2015},
abstract={This paper aims at outlining some aspects of biolinguistics — the study of the biological bases of language. After a brief introduction, some considerations are made about the reasons for the lack of attention given to this approach until very recently, as well as some reasons for its reemergence, which is now underway.},
booktitle={Textos Selecionados do XXX Encontro Nacional da Associação Portuguesa de Linguística},
pages = {381--401},
title= {A reemergência da Biolinguística},
address = {Porto},
publisher = {FLUP/APL}}

@incollection{boeckxmartins16,
title = {Biolinguistics},
author = {Boeckx, Cedric and Martins, Pedro Tiago},
year = {2016},
editor = {Aronoff, Mark},
booktitle = {Oxford Research Encyclopedia of Linguistics},
doi = {10.1093/acrefore/9780199384655.013.20},
abstract = {All humans can acquire at least one natural language. Biolinguistics is the name given to the interdisciplinary enterprise that aims to unveil the biological bases of this unique capacity.}
}

@article{boeckx2017conjecture,
title={A conjecture about the neural basis of recursion in light of descent with modification},
author={Boeckx, Cedric},
journal={{Journal of Neurolinguistics}},
volume={43},
pages={193--198},
year={2017},
publisher={Elsevier},
doi={10.1016/j.jneuroling.2016.08.003},
abstract={The goal of this paper is to examine the possible neurobiological basis of a defining property of the human language faculty: recursion. I suggest that recursion should be understood in light of Darwinian's descent with modification. Descent: that is, based on ingredients of neural circuitry found in ‘non-linguistic’ species; and modification: a reconfiguration that is specific to anatomically modern humans. I argue that the expansion of the parietal region associated with the globularization of the neurocranium in our species contributed to the transformation of the connection between Broca's and Wernicke's region via Geschwind's territory, and enabled the pairing of evolutionary ancient networks that together became capable of constructing and processing not just sequences, but sequences of sequences.}
}

@article{boeckx2014functional,
title={The functional neuroanatomy of serial order in language},
author={Boeckx, Cedric and Martinez-Alvarez, Anna and Leivada, Evelina},
journal={{Journal of Neurolinguistics}},
volume={32},
pages={1--15},
year={2014},
publisher={Elsevier},
doi={10.1016/j.jneuroling.2014.07.001},
abtract={The paper aims to shed light on how serial order is computed in the human mind/brain, focusing on the nature of linearization in language. Linearization is here understood as the mapping of hierarchical syntactic structures onto linear strings. We take as our point of departure the now well-established need to subdivide Broca's region into different areas, and claim that these brain areas play important and distinct roles in the context of linearization. Crucially, for this mapping to be valid, linearization must be decomposed into a series of distinct (generic) sub-operations. Thus, the present work highlights the benefit of decomposing Broca's area and the linearization algorithm in parallel to formulate linking hypotheses between mind and brain.,}
}

@ARTICLE{martinsboeckx16,
Author={Martins, Pedro Tiago and Boeckx, Cedric},
Title={What we talk about when we talk about biolinguistics},
Year={2016},
Abstract={The study of the biological foundations of language is sometimes called biolinguistics. This particular term finds its historical origins in the 1950s, and for various reasons it has also gained considerable traction in recent years. While its increasing use apparently signals an equally increasing interest in biology, apart from a few exceptions not much is added to and beyond standard linguistic theorizing by those linguists who use it, resulting in a complex and confusing literature. This state of affairs has led, on the one hand, to the perpetuation of biologically implausible proposals that have pervaded linguistics for decades, and to ill-placed criticism on the progress and even the very legitimacy of a biologically-informed study of language, on the other. By reviewing different ways in which research under the biolinguistics moniker has been carried out, as well as some common criticisms, we hope to dispel some misconceptions about what constitutes a biolinguistic approach, as well as point out what we contend is real progress in the study of the biological bases and evolution of the human language faculty, to which the term is better and rightly applied.},
journal = {Linguistics Vanguard},
number = {1},
volume = {2},
doi = {10.1515/lingvan-2016-0007}
}

@incollection{martins17,
author = {Martins, Pedro Tiago},
year = {2017},
title = {There is no place for markedness in biologically-informed phonology},
booktitle = {Beyond markedness in formal phonology},
editor = {Samuels, Bridget D},
publisher = {John Benjamins},
address = {Amsterdam},
pages = {219--232},
url = {https://benjamins.com/#catalog/books/la.241/main},
abstract = {Markedness is a pervasive notion in theories of language. In some phonological theories or models, it very much occupies a prominent position. In this chapter it is argued that the notion or markedness is not useful to our understanding of phonology and language in general, for two reasons. The first is that the notion of markedness has convincingly been shown in the literature to be a confusing label for a variety of things, all of which can be explained independently. The second is that if phonology is to be understood as part of a biological system (which we call language), its components must be amenable to investigation in the cognitive and biological sciences. The notion of markedness as part of a phonological component does not seem to qualify for any meaningful kind of investigation in that respect. It follows that markedness has no place in biologically-informed phonology.}
}

@article{alamri_etal2015,
author={Alamri, Saleh and Zhang, Elizabeth Q and Theofanopoulou, Constantina and Castillo, Gonzalo and Shi, Edward R and Martins, Pedro Tiago and Martínez Ferreiro, Silvia and Boeckx, Cedric},
title={Implications of Subcortical structures in Aphasia},
journal={{Frontiers in Psychology}},
volume={81},
year={2015},
url={https://www.frontiersin.org/10.3389/conf.fpsyg.2015.65.00081/event_abstract},
doi={10.3389/conf.fpsyg.2015.65.00081},
issn={1664-1078}
}

@article{theofanopoulou2015cognitive,
title={Cognitive phylogenies, the Darwinian logic of descent, and the inadequacy of cladistic thinking},
author={Theofanopoulou, Constantina and Boeckx, Cedric},
journal={{Frontiers in Cell and Developmental Biology}},
doi = {https://doi.org/10.3389/fcell.2015.00064},
volume={3},
year={2015}
}

@article{theofanopoulou_etal17,
author = {Theofanopoulou, Constantina and Gastaldon, Simone and O'Rourke, Thomas and Samuels, Bridget D. and Messner, Angela and Martins, Pedro Tiago and Delogu, Francesco and Alamri, Saleh and Boeckx, Cedric},
title = {Self-domestication in Homo sapiens: Insights from comparative genomics},
year = {2017},
volume = {12},
number = {10},
pages = {e0185306},
doi = {10.1371/journal.pone.0185306},
abstract = {This study identifies and analyzes statistically significant overlaps between selective sweep screens in anatomically modern humans and several domesticated species. The results obtained suggest that (paleo-)genomic data can be exploited to complement the fossil record and support the idea of self-domestication in Homo sapiens, a process that likely intensified as our species populated its niche. Our analysis lends support to attempts to cap- ture the “domestication syndrome” in terms of alterations to certain signaling pathways and cell lineages, such as the neural crest.},
URL = {http://journals.plos.org/plosone/article/file?id=10.1371/journal.pone.0185306&type=printable},
journal = {{PLoS ONE}},
press = {How humans (maybe) domesticated themselves. ScienceNews 191(13): 26. July 8, 2017}
}

@article{boeckx2017,
author={Boeckx, Cedric},
title={The language-ready head: Evolutionary considerations},
journal={Psychonomic Bulletin {\&} Review},
year={2017},
volume={24},
number={1},
pages={194--199},
abstract={This article offers a succinct overview of the hypothesis that the evolution of cognition could benefit from a close examination of brain changes reflected in the shape of the neurocranium. I provide both neurological and genetic evidence in support of this hypothesis, and conclude that the study of language evolution need not be regarded as a mystery.},
doi={10.3758/s13423-016-1087-5},
url={https://doi.org/10.3758/s13423-016-1087-5}
}

@article{boeckx2016,
author={Boeckx, Cedric and Benítez-Burraco, Antonio},
title={Editorial: Components of the Language-Ready Brain},
journal={{Frontiers in Psychology}},
volume={7},
pages={762},
year={2016},
url={http://journal.frontiersin.org/article/10.3389/fpsyg.2016.00762},
doi={10.3389/fpsyg.2016.00762}
}

@article{boeckx_benitez2015,
author={Boeckx, Cedric and Benítez-Burraco, Antonio},
title={Osteogenesis and neurogenesis: a robust link also for language evolution},
journal={{Frontiers in Cellular Neuroscience}},
volume={9},
pages={291},
year={2015},
url={http://journal.frontiersin.org/article/10.3389/fncel.2015.00291},
doi={10.3389/fncel.2015.00291}
}

@article{asano_boeckx_2015,
author={Asano, Rie and Boeckx, Cedric},
title={Syntax in language and music: what is the right level of comparison?},
journal={{Frontiers in Psychology}},
journal={6},
pages={942},
year={2015},
url={http://journal.frontiersin.org/article/10.3389/fpsyg.2015.00942},
doi={10.3389/fpsyg.2015.00942},
abstract={It is often claimed that music and language share a process of hierarchical structure building, a mental “syntax”. Although several lines of research point to commonalities, and possibly a shared syntactic component, differences between “language syntax” and “music syntax” can also be found at several levels: conveyed meaning, and the atoms of combination, for example. To bring music and language closer to one another, some researchers have suggested a comparison between music and phonology (‘phonological syntax’), but here too, one quickly arrives at a situation of intriguing similarities and obvious differences. In this paper we suggest that a fruitful comparison between the two domains could benefit from taking the grammar of action into account. In particular, we suggest that what is called “syntax” can be investigated in terms of goal of action, action planning, motor control, and sensory-motor integration. At this level of comparison, we suggest that some of the differences between language and music could be explained in terms of different goals reflected in the hierarchical structures of action planning: the hierarchical structures of music arise to achieve goals with a strong relation to the affective-gestural system encoding tension-relaxation patterns as well as socio-intentional system, whereas hierarchical structures in language are embedded in a conceptual system that gives rise to compositional meaning. Similarities between music and language are most clear in the way several hierarchical plans for executing action are processed in time and sequentially integrated to achieve various goals.}
}

@article{benitez_boeckx2015,
author={Benítez-Burraco, Antonio and Boeckx, Cedric},
title={Possible functional links among brain- and skull-related genes selected in modern humans},
journal={{Frontiers in Psychology}},
volume={6},
pages={794},
year={2015},
url={http://journal.frontiersin.org/article/10.3389/fpsyg.2015.00794},
doi={10.3389/fpsyg.2015.00794},
abstract={The sequencing of the genomes from extinct hominins has revealed that changes in some brain-related genes have been selected after the split between anatomically-modern humans and Neanderthals/Denisovans. To date, no coherent view of these changes has been provided. Following a line of research we initiated in Boeckx and Benítez-Burraco (2014a), we hypothesize functional links among most of these genes and their products, based on the existing literature for each of the gene discussed. The genes we focus on are found mutated in different cognitive disorders affecting modern populations and their products are involved in skull and brain morphology, and neural connectivity. If our hypothesis turns out to be on the right track, it means that the changes affecting most of these proteins resulted in a more globular brain and ultimately brought about modern cognition, with its characteristic generativity and capacity to form and exploit cross-modular concepts, properties most clearly manifested in language.}
}

@article{boeckx_theo2015,
author={Boeckx, Cedric and Theofanopoulou, Constantina},
title={Commentary on: Labels, cognomes, and cyclic computation: an ethological perspective},
journal={{Frontiers in Psychology}},
volume={6},
pages={784},
year={2015},
url={http://journal.frontiersin.org/article/10.3389/fpsyg.2015.00784},
doi={10.3389/fpsyg.2015.00784}
}

@article{benitez_boeckx2015approaching,
author={Benítez-Burraco, Antonio and Boeckx, Cedric},
title={Approaching motor and language deficits in autism from below: a biolinguistic perspective},
journal={{Frontiers in Integrative Neuroscience}},
volume={9},
pages={25},
year={2015},
url={http://journal.frontiersin.org/article/10.3389/fnint.2015.00025},
doi={10.3389/fnint.2015.00025},
issn={1662-5145}
}

@article{martins_boeckx2014,
author={Martins, Pedro T. and Boeckx, Cedric},
title={Attention mechanisms and the mosaic evolution of speech},
journal={{Frontiers in Psychology}},
volume={5},
pages={1463},
year={2014},
url={http://journal.frontiersin.org/article/10.3389/fpsyg.2014.01463},
doi={10.3389/fpsyg.2014.01463},
issn={1664-1078},
abstract={There is still no categorical answer for why humans, and no other species, have speech, or why speech is the way it is. Several purely anatomical arguments have been put forward, but they have been shown to be false, biologically implausible, or of limited scope. This perspective paper supports the idea that evolutionary theories of speech could benefit from a focus on the cognitive mechanisms that make speech possible, for which antecedents in evolutionary history and brain correlates can be found. This type of approach is part of a very recent, but rapidly growing tradition, which has provided crucial insights on the nature of human speech by focusing on the biological bases of vocal learning. Here, we call attention to what might be an important ingredient for speech. We contend that a general mechanism of attention, which manifests itself not only in visual but also auditory (and possibly other) modalities, might be one of the key pieces of human speech, in addition to the mechanisms underlying vocal learning, and the pairing of facial gestures with vocalic units.}
}

@article{leivada_boeckx2014,
author={Leivada, Evelina and Boeckx, Cedric},
title={Schizophrenia and cortical blindness: protective effects and implications for language},
journal={{Frontiers in Human Neuroscience}},
volume={8},
pages={940},
year={2014},
url={http://journal.frontiersin.org/article/10.3389/fnhum.2014.00940},
doi={10.3389/fnhum.2014.00940},
abstract={The repeatedly noted absence of case-reports of individuals with schizophrenia and congenital/early developed blindness has led several authors to argue that the latter can confer protective effects against the former. In this work, we present a number of relevant case-reports from different syndromes that show comorbidity of congenital and early blindness with schizophrenia. On the basis of these reports, we argue that a distinction between different types of blindness in terms of the origin of the visual deficit, cortical or peripheral, is crucial for understanding the observed patterns of comorbidity. We discuss the genetic underpinnings and the brain structures involved in schizophrenia and blindness, with insights from language processing, laying emphasis on the three structures that particularly stand out: the occipital cortex, the lateral geniculate nucleus and the pulvinar. Last, we build on previous literature on the nature of the protective effects in order to offer novel insights into the nature of the protection mechanism from the perspective of the brain structures involved in each type of blindness.}
}

@article{benitez2016globularization,
title={Globularization and domestication},
author={Benítez-Burraco, Antonio and Theofanopoulou, Constantina and Boeckx, Cedric},
journal={Topoi},
pages={265--278},
volume = {37},
number = {2},
year={2016},
publisher={Springer},
doi = {10.1007/s11245-016-9399-7},
abstract = {This paper aims to explore a potential connection between two hypotheses recently put forward in the context of language evolution. One hypothesis argues that some human-specific change(s) in the hominin brain developmental program habilitated the neuronal workspace that enabled “cognitive modernity” to unfold, also resulting in our globularized braincase. The other argues that the cultural niche resulting from our self-domestication favored the emergence of natural languages. In this article we document numerous links between the genetic changes we have claimed may have brought about globularization and neural crest cells, which have been claimed to explain the constellation of distinctive traits (physical, cognitive, and behavioral) found in all domesticated mammals. If these links turn out to be as robust as we think they are, globularization and self-domestication may well be closely related phenomena in the context of human evolution.}
}

@article{boeckx_benitez2014,
author={Boeckx, Cedric and Benítez-Burraco, Antonio},
title={Globularity and language-readiness: generating new predictions by expanding the set of genes of interest},
journal={{Frontiers in Psychology}},
volume={5},
pages={1324},
year={2014},
url={http://journal.frontiersin.org/article/10.3389/fpsyg.2014.01324},
doi={10.3389/fpsyg.2014.01324},
abstract={This study builds on the hypothesis put forth in Boeckx and Benitez-Burraco (2014), according to which the developmental changes expressed at the levels of brain morphology and neural connectivity that resulted in a more globular braincase in our species were crucial to understand the origins of our language-ready brain. Specifically, this paper explores the links between two well-known ‘language-related’ genes like FOXP2 and ROBO1 implicated in vocal learning and the initial set of genes of interest put forth in Boeckx and Benitez-Burraco (2014), with RUNX2 as focal point. Relying on the existing literature, we uncover potential molecular links that could be of interest to future experimental inquiries into the biological foundations of language and the testing of our initial hypothesis. Our discussion could also be relevant for clinical linguistics and for the interpretation of results from paleogenomics.}
}

@article{benitez_boeckx2014,
author={Benítez-Burraco, Antonio and Boeckx, Cedric},
title={FOXP2, retinoic acid, and language: a promising direction},
journal={{Frontiers in Cellular Neuroscience}},
volume={8},
pages={387},
year={2014},
url={http://journal.frontiersin.org/article/10.3389/fncel.2014.00387},
doi={10.3389/fncel.2014.00387}
}

@article{benitez_boeckx2014revneurol,
author={Benítez-Burraco, Antonio and Boeckx, Cedric},
title = {Trastornos del lenguaje como oscilopatías},
journal = {{Revista de Neurologia}},
volume = {59},
number = {7},
pages = {333--334},
year={2014},
url = {https://www.neurologia.com/articulo/2014317},
}

@article{boeckx_fujita2014,
author={Boeckx, Cedric and Fujita, Koji},
title={Syntax, action, comparative cognitive science, and Darwinian thinking},
journal={{Frontiers in Psychology}},
volume={5},
pages={627},
year={2014},
url={http://journal.frontiersin.org/article/10.3389/fpsyg.2014.00627},
doi={10.3389/fpsyg.2014.00627},
issn={1664-1078}
}

@article{benitez_boeckx2014evodevo,
title={Universal Grammar and biological variation: an EvoDevo agenda for comparative biolinguistics},
author={Benítez-Burraco, Antonio and Boeckx, Cedric},
journal={{Biological theory}},
volume={9},
number={2},
pages={122--134},
year={2014},
url={https://link.springer.com/article/10.1007%2Fs13752-014-0164-0},
doi={10.1007/s13752-014-0164-0},
issn={1555-5550}
}

@article{10.3389/fpsyg.2014.00282,
author={Boeckx, Cedric and Benítez-Burraco, Antonio},
title={The shape of the human language-ready brain},
journal={{Frontiers in Psychology}},
volume={5},
pages={282},
year={2014},
url={http://journal.frontiersin.org/article/10.3389/fpsyg.2014.00282},
doi={10.3389/fpsyg.2014.00282},
abstract={Our core hypothesis is that the emergence of our species-specific language-ready brain ought to be understood in light of the developmental changes expressed at the levels of brain morphology and neural connectivity that occurred in our species after the split from Neanderthals-Denisovans and that gave us a more globular braincase configuration. In addition to changes at the cortical level, we hypothesize that the anatomical shift that led to globularity also entailed significant changes at the subcortical level. We claim that the functional consequences of such changes must also be taken into account to gain a fuller understanding of our linguistic capacity. Here we focus on the thalamus, which we argue is central to language and human cognition, as it modulates fronto-parietal activity. With this new neurobiological perspective in place, we examine its possible molecular basis. We construct a candidate gene set whose members are involved in the development and connectivity of the thalamus, in the evolution of the human head, and are known to give rise to language-associated cognitive disorders. We submit that the new gene candidate set opens up new windows into our understanding of the genetic basis of our linguistic capacity. Thus, our hypothesis aims at generating new testing grounds concerning core aspects of language ontogeny and phylogeny.}
}

@article{bruner_boeckx2013,
author={Bruner, Emiliano and Boeckx, Cedric},
title = {Language: the elusive milestone},
journal = {{Journal of Anthropological Sciences}},
volume = {91},
year = {2013},
pages = {13--24},
url = {http://www.isita-org.com/jass/Contents/2013vol91/Bruner/24371148.pdf},
doi = {10.4436/JASS.91025},
issn = {1827-4765}
}

@article{boeckx2013,
author={Boeckx, Cedric},
title = {Biolinguistics: forays into human cognitive biology},
journal = {{Journal of Anthropological Sciences}},
volume = {91},
year = {2013},
pages = {63--89},
url = {http://www.isita-org.com/jass/Contents/2013vol91/Boeckx/24038628.pdf},
doi = {10.4436/jass.91009},
abstract = {The present article surveys the field of biolinguistics. It revisits the roots of the field’s core research agenda, then turns to the various factors that led to its recent re-emergence, and finally offers suggestions for future inquiry. The essay also serves to highlight certain conceptual issues that should be addressed if the field is to bear its interdisciplinary fruits.}
}

@article{boeckx_leivada2013,
author = {Boeckx, Cedric AND Leivada, Evelina},
journal = {{PLoS ONE}},
publisher = {Public Library of Science},
title = {Entangled Parametric Hierarchies: Problems for an Overspecified Universal Grammar},
year = {2013},
volume = {8},
url = {https://doi.org/10.1371/journal.pone.0072357},
pages = {1-11},
abstract = {This study addresses the feasibility of the classical notion of parameter in linguistic theory from the perspective of parametric hierarchies. A novel program-based analysis is implemented in order to show certain empirical problems related to these hierarchies. The program was developed on the basis of an enriched data base spanning 23 contemporary and 5 ancient languages. The empirical issues uncovered cast doubt on classical parametric models of language acquisition as well as on the conceptualization of an overspecified Universal Grammar that has parameters among its primitives. Pinpointing these issues leads to the proposal that (i) the (bio)logical problem of language acquisition does not amount to a process of triggering innately pre-wired values of parameters and (ii) it paves the way for viewing language, epigenetic (‘parametric’) variation as an externalization-related epiphenomenon, whose learning component may be more important than what sometimes is assumed.},
number = {9},
doi = {10.1371/journal.pone.0072357}
}